See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/312217513

New species of ptyctimous (, ) from the Caribbean

Article in Systematic and Applied Acarology · January 2017 DOI: 10.11158/saa.22.2.8

CITATIONS READS 0 90

2 authors:

Wojciech Niedbała Sergey G. Ermilov Adam Mickiewicz University Tyumen State University

256 PUBLICATIONS 727 CITATIONS 388 PUBLICATIONS 1,076 CITATIONS

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Taxonomy, biodiversity and ecology of oribatid mites (Acari, Oribatida) of New Zealand View project

Taxonomy, biodiversity and ecology of oribatid mites (Acari, Oribatida) of Vietnam View project

All content following this page was uploaded by Sergey G. Ermilov on 24 January 2017.

The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document and are linked to publications on ResearchGate, letting you access and read them immediately. Systematic & Applied Acarology 22(2): 241–252 (2017) ISSN 1362-1971 (print) http://doi.org/10.11158/saa.22.2.8 ISSN 2056-6069 (online)

Article http://zoobank.org/urn:lsid:zoobank.org:pub:04E3FFFA-4720-48C9-84C0-63AEA762ED25 New species of ptyctimous mites (Acari, Oribatida) from the Caribbean

WOJCIECH NIEDBAŁA1 & SERGEY G. ERMILOV2* 1Faculty of Biology, Department of Taxonomy and Ecology, Adam Mickiewicz University, Poznan, Poland. E-mail: [email protected] 2Tyumen State University, Tyumen, Russia. E-mail: [email protected] *Corresponding author

Abstract

A list of identified oribatid ptyctimous mites from Lesser Antilles and Bahamas, including 18 species, six genera and three families, is provided. Of these, three species are new records for the Caribbean, and three species are new to science. Oribotritia grossa Niedbała sp. nov. differs from other neotropical Oribotritia by the presence of relatively short, thick, distally blunt sensilli. Arphthiracarus alienus Niedbała sp. nov. differs from other Arphthiracarus species by the presence of uniquely shaped sensilli, short spiniform setae, the median sigillar field long and narrow with an incision between the rostral setae, four pairs of notogastral lyrifissures and the location of vestigial setae f1 anterior to h1. Arphthiracarus strictus Niedbała sp. nov. differs from A. brasiliensis Niedbała, 1988, A. iubatus Niedbała, 2003, A. latebrosus (Niedbała, 1982) and A. pervalidus Niedbała, 2003 by the presence of a unique tight location of genital setae g4-6 and bifurcated setae d of femora I. In addition, descriptions of three tritonymphs are presented.

Key words: ptyctimous mites, fauna, list of taxa, new species, morphology, systematics, ontogeny, tritonymph, Neotropical region

Introduction

Ptyctimous mites (Acari, Oribatida) of the Caribbean are insufficiently known (e.g. Mahunka 1985, 1998; Niedbała 2004, 2009; Niedbała & Ermilov 2015). Until now, 73 species are known: 39 species of Phthiracaroidea, 28 species of Euphthiracaroidea and six species of Arthroptyctima. Nearly half of these species (31 species) were found in Cuba. Our investigation is based on material collected from the Lesser Antilles and Bahamas by Dr. P.W. Hummelinck in 1936–1967 during his voyages in the Neotropical region. In the course of taxonomic identification we discovered three new species, one species belonging to the genus Oribotritia Jacot, 1924 () and two to Arphthicarus Niedbała, 1994 (Steganacaridae). The main goal of the paper is to describe and illustrate these new species and to present a list of identified ptyctimous taxa. The generic characters of Oribotritia and Arphthicarus, and identification keys and analysis on geographical distribution to the Neotropical species were presented by Niedbała (2004). In addition, descriptions of three tritonymphs, belonging most likely to Indotritia (Indotritia) krakatauensis (Sellnick, 1923) and Arphthicarus alienus Niedbała sp. nov. are presented. Our identification of juveniles is based on the following reasons: 1) of the oribatid mites in the samples, only specimens of this species were present; 2) adults were of the appropriate size to have tritonymphs with such dimensions.

© Systematic & Applied Acarology Society 241 At present, the juveniles of ptyctimous mites are poorly known (Norton & Ermilov 2014), therefore despite approximate species identification of tritonymphs and few specimens, we included their descriptions in this paper.

Material and methods

Material. Ptyctimous mites were collected by Dr. P.W. Hummelinck (and sorted by Dr. M. Sellnick) from litter/soil in the following sites: 295 (this code–hereinafter–to link specimens with data in the Naturalis Biodiversity Center, Leiden, The Netherlands)—Trinidad, Tetron Bay, NW Trinidad, 7.V.1936; 298A—Saba, road to the Bottom at S-curve, 17°37'N, 63°14'W, 18.III.1937; 391—Antigua, near Bat’s Cave, East of Nelson’s Dockyard, limestone, considerable growth of xerophytic shrubs, below rocks and in fissures with some plant decay, 17°1'0"N, 61°45'0" W, 13.VII.1955; 420—St. Kitts and Nevis, St Kitts, Wingfield River, 30.VI.1949; 424—St. Eustatius, White Wall base, 17.469555°N, 62.956178°W, 109 m a.s.l., 6.VIII.1949; 451—Saint-Barthelémy, Public, 17°54'6.47"N, 62°51'9.27"W, 4.VI.1949; 482—Anguilla, Forest Point, 18.VI.1949; 483—Anguilla, Long Bay, 18°11'27.6"N, 63°07'53"W, 18.VI.1949; 486—Anguilla, Upper Prickley Pear Island, 18°15'50.20"N, 63°10'26.62"W, 17.VI.1949; 493—Bahamas, New Providence Island, Blue Hills at Hunt's Cave, 22.VIII.1949; 572—Trinidad, N-range, Cerro del Aripo, 10°44'N, 61°15'W, 30.I.1955; 576—Trinidad, Gaspar Grande Island, chimney of cave, 10°40'N, 61°38'W, 11.I.1955; 580—Trinidad, Chacachacare Island, Bande du Sud, 10°40'30.31"N, 61°45'26.24"W, 1.I.1955; 585—Tobago, Little Tobago Island, at landing, volcanic rock, shrubs and scattered small trees, rock fissures with debris and decay, 11°18'2.02"N, 60°30'18.52"W, 18.I.1955; 588—Grenada, Corinth Estate, St. David, volcanic rock, cocoa estate with banana and coconut trees, leaf decay of Theobroma, Musa and Cocos, 25.I.1955; 591—Antigua, near Bats Cave, 17°1'0"N, 61°45'0"W, 13.VII.1955; 595A—Antigua, near Yepton Mill, W of St. John's, decomposed rocks, cultivated area with grasses, shrubs, cacti and Tamarindus, under pieces of rock, dead wood, 17°7'N, 61°52'W, 17.VII.1955; 596—Barbuda, Martello Tower, 17º35'38.73"N, 61º49'49.40"W, 8.VII.1955; 632—Aruba, Mangel Corá tunnel, Lago Oil Company refinery, 12.4203ºN, 69.8694ºW, 2.V.1955; 714—Saba, Cove Bay, andesite, shore vegetation, sandy leaf decay of Coccoloba uvifera, 17°38'34.43"N, 63°13'17.74"W, 5.X.1963; 829—Sint Maarten, Point Blanche Bay, 18°0'44.63"N, 63°2'11.05"W, 9.VII.1967. Methods. The mites were submerged in lactic acid and mounted on temporary cavity slides for the duration of the study. Body measurements are presented in micrometers. Length of the body setae was measured in lateral aspect. Holotypes of new species are deposited in the collection of the Netherlands Centre for Biodiversity Naturalis, Leiden, The Netherlands; paratypes are deposited in the collection of the Department of Animal Taxonomy and Ecology, Adam Mickiewicz University, Pozna , Poland. In the List of identified ptyctimous mites subsection, the authors and years of species are not included in the References section.

242 SYSTEMATIC & APPLIED ACAROLOGY VOL. 22 List of identified ptyctimous mites

Localities (as in Material section) given after the species name and number of individuals in parentheses.

Oribotritiidae —Oribotritia grossa Niedbała sp. nov.: 585 (28) —Mesotritia multisetosa Niedbała, 2004: 451 (12), 482 (13), 486 (2), 572 (1), 591 (1), 596 (1) (new record for the Caribbean) —Indotritia (Indotritia) bellingeri Niedbała & Schatz, 1996: 420 (1), 576 (2), 588 (1) —Indotritia (Indotritia) krakatauensis (Sellnick, 1923): 391 (7), 424 (2), 451 (8), 482 (6), 483 (4), 486 (1), 493 (1), 572 (1), 576 (1), 580 (3), 588 (12), 591 (1), 595A (5), 596 (25), 632 (1), 714 (6) —Indotritia (Indotritia) retusa Niedbała & Schatz, 1996: 298A (1), 424 (2), 451 (8), 572 (3), 588 (2), 596 (3)

Euphthiracaridae —Acrotritia ardua (Koch, 1841): 576 (2) —Acrotritia clavata (Märkel, 1964): 420 (1), 424 (1), 482 (2), 486 (2), 576 (2), 591 (1) —Acrotritia dikra (Niedbała & Schatz, 1996): 295 (1), 486 (1), 493 (1) —Acrotritia vestita (Berlese, 1913): 451 (2), 493 (1), 572 (1), 588 (1)

Steganacaridae —Arphthicarus alienus Niedbała sp. nov.: 295 (5), 576 (7) —Arphthicarus carinatus Niedbała, 2004: 572 (1) —Arphthicarus latebrosus (Niedbała, 1982): 451 (1), 576 (2), 585 (4) —Arphthicarus parasaucius Niedbała, 2003: 576 (4), 585 (11) —Arphthicarus saucius (Niedbała, 1988): 572 (1) (new record for the Caribbean) —Arphthicarus strictus Niedbała sp. nov.: 572 (1) —Atropacarus (Hoplophorella) hamatus (Ewing, 1909): 420 (1), 595A (1) —Atropacarus (Hoplophorella) lanceosetus (Balogh & Mahunka, 1981): 829 (1) (new record for the Carib- bean) —Atropacarus (Hoplophorella) vitrinus (Berlese, 1913): 572 (1)

Descriptions of new taxa

Oribotritia grossa Niedbałasp. nov. (Fig. 1A–F)

Diagnosis. Sensilli of prodorsum relatively short, thick rigid, smooth, blunt distally; interlamellar, lamellar and rostral setae short, fine, procumbent; setae c1 of notogaster remote from anterior border, setae c2 and c3 situated near border; aggenital setae ag1 considerably shorter than ag2; two pairs of anal and two pairs of adanal setae. Description. Measurements. Medium to big sized species. Holotype. Prodorsum: length 454, width 343, height 156; prodorsal setae: ss 73, in 61, le 51, ro 58; notogaster: length 858, width and height 656; notogastral setae: c1 71, h1, ps1 78, c3 76; genito-aggenital plate 202 × 116; ano-adanal plate 429 × 86; aggenital setae: ag1 25, ag2 53. Paratypes. Prodorsum: length 379–945, height 116– 192; notogaster: length 697–1141, height 454–858. Integument. Colour brown. Integument very finely pitted. Prodorsum with simple, long and thick lateral carinae. Sensilli (ss) relatively short, thick rigid, smooth, blunt distally. Interlamellar (in), lamellar (le) and rostral (ro) setae short, fine, procumbent, exobothridial (ex) setae vestigial; relation of lengths: ss>in>ro>le.

2017 NIEDBAŁA& ERMILOV: NEW SPECIES OF PTYCTIMOUS MITES FROM THE CARIBBEAN 243 FIGURE 1. Oribotritia grossa Niedbała sp. nov. (holotype): A—prodorsum, dorsal view; B—prodorsum, lateral view; C—opisthosoma, lateral view; D—mentum of subcapitulum; E—right side of ventral plates; F— trochanter and femur of leg I. Scale bars 100 μm (A–C, E), 50 μm (D, F).

Notogaster with setae short, fine, rough, setae c1 remote from anterior border, setae c2 and c3 situated near border. Ventral region. Seta h of mentum considerably longer than distance between them. Nine pairs of genital setae, setae g1-5 in progenital position remote from genital setae g6-9. Two pairs of aggenital setae present, ag1 considerably shorter than ag2. Two pairs of anal and two pairs of adanal setae present, ad1 located lateral and posterior to an1. Lyrifissures iad located lateral and posterior to ad2.

244 SYSTEMATIC & APPLIED ACAROLOGY VOL. 22 Legs. Chaetome (without tarsi): I: 1-4-5(2)-5(1), II: 1-4-4(1)-4(1), III: 3-2-3(1)-3(1), IV: 3-2- 2(1)-3(1). Femora I with short distal spine. Material examined. Holotype and 27 paratypes: Antilles, Tobago, Little Tobago Island, at landing, volcanic rock, shrubs and scattered small trees, rock fissures with debris and decay, 11°18'2.02"N, 60°30'18.52"W, litter and soil, 18.I.1955 (P.W. Hummelinck). Etymology. The name of the new species grossa (masc. grossus)is Latin for “thick” and alludes to the thickened shape of the sensilli, which is unique in this genus. Remarks. Oribotritia grossa sp. nov. differs from all neotropical Oribotritia species by the presence of relatively short, thick, distally blunt sensilli. All neotropical species has sensilli pointed distally. Apart from the sensilli, only one neotropical species, O. didyma Niedbała & Schatz, 1996, is similar to the new species by the presence of two pairs of aggenital setae, two pairs of anal and two pairs of adanal setae, but differs by the presence of two pairs of prodorsal carinae (versus one pair in O. grossa sp. nov.) and adanal setae ad1 located lateral and anterior from anal setae an1 (versus located posterior in O. grossa sp. nov.).

Arphthicarus alienus Niedbałasp. nov. (Fig. 2A–I)

Diagnosis. Sensilli short, clavate, inside fusiform with extended apical end, other setae short spiniform; four pairs of notogastral lyrifissures. Description. Measurements. Species of rather big size. Holotype. Prodorsum: length 419, width 278, height 177; prodorsal setae: ss 56, in and le 30, ro 76, ex 53; notogaster: length 778, width 565, height 515; notogastral setae: c1, d1 78, c2, h1, ps1 71, c3 32, c1/c1-d1=0.5; genito-aggenital plate 222 × 162; ano-adanal plate 313 × 156. Paratypes. Prodorsum: length 395–495, height 182–222; notogaster: length 727–929, height 501–697. Integument. Colour brown. Surface of body with weak concavities. Prodorsum with weak median crista. Lateral carinae absent. Sigillar fields long and narrow, median longer than laterals with incision between the rostral setae. Posterior furrows distinct. Sensilli with unique shape, rather short, clavate, inside fusiform with extended apical end. Interlamellar, lamellar and rostral setae spiniform, rough; relation of lengths: ro>ss>ex>in=le.

Notogaster with 15 pairs of short, spiniform setae, slightly serrate at distal end. Setae c1 and c3 considerably remote from anterior margin, c2 more remote from margin, c3 smallest. Vestigial setae f1 anterior to h1. All four pairs of lyrifissures ia, im, ip and ips present. Ventral region. Setae h of mentum considerably longer than distance between them. Arrangement of genital setae: 6(4+2): 3. An-oadanal plates with five pairs of short, spiniform setae, ad2 longer than ad1, anal setae and ad3 the shortest. Legs. Chaetome of legs of “complete type”. Setae d on femora remote from distal end of article; setae v” very long, setae v’ minute. Material examined. Holotype and 6 paratypes: Antilles, Trinidad, Gaspar Grande Island, chimney of cave, 10°40'N, 61°38'W, litter and soil, 11.I.1955 (P.W. Hummelinck). Five paratypes: Antilles, Trinidad, Tetron Bay, NW Trinidad, litter and soil, 7.V.1936 (P.W. Hummelinck). Etymology. The name of the new species alienus is Latin for “another, strange, foreign, unrelated” and alludes to the unique construction of sensilli. Remarks. The new species is easily distinguishable from congeners by the presence of the uniquely shaped sensilli, short spiniform setae, median sigillar field long and narrow with incision between the rostral setae, four pairs of notogastral lyrifissures and the location of vestigial setae f1 anterior to h1.

2017 NIEDBAŁA& ERMILOV: NEW SPECIES OF PTYCTIMOUS MITES FROM THE CARIBBEAN 245 FIGURE 2. Arphthicarus alienus Niedbała sp. nov. (holotype): A—prodorsum, dorsal view; B—sensillus, dorsal view; C—prodorsum, lateral view; D—lateral view of opisthosoma; E—seta c1; F—mentum of subcapitulum, G—right genito-aggenital plate, H—right ano-adanal plate, I—trochanter and femur of leg I. Scale bars 100 μm (A, C, D, G, H), 50 μm (F, I), 25 μm (B, E).

246 SYSTEMATIC & APPLIED ACAROLOGY VOL. 22 Arphthicarus strictus Niedbałasp. nov. (Fig. 3A–I)

Diagnosis. Tight location of genital setae g4-6; bifurcated setae d of femora I; sensilli short, with narrow pedicel and rounded head; interlamellar, lamellar and rostral setae short, spiniform; notogastral setae rather short, covered with spines, obtuse distally; vestigial setae f1 situated at the level of h1 setae; four pairs of lyrifissures.

FIGURE 3. Arphthicarus strictus Niedbała sp. nov. (holotype): A—prodorsum, dorsal view; B—prodorsum, lateral view; C—lateral view of opisthosoma; D—mentum of subcapitulum, E—genito-aggenital plates, F— position of setae g4-6 of genito-aggenital plates; G—left ano-adanal plate, H—trochanter and femur of leg I, I— seta d of femora I. Scale bars 100 μm (A–C, E, G), 50 μm (D, F, H), 25 μm (I).

2017 NIEDBAŁA& ERMILOV: NEW SPECIES OF PTYCTIMOUS MITES FROM THE CARIBBEAN 247 Description. Measurements. Species of rather big size. Holotype. Prodorsum: length 364, width 245, height 136; prodorsal setae: ss 51, in 56, le 25, ro 61; notogaster: length 757, width 525, height

485; notogastral setae: c1 136, h1 94, ps1 147, c1/c1-d1 = 0.8; genito-aggenital plate 242 × 116; ano- adanal plate 262 × 116. Integument. Colour light brown. Surface of body covered with weak concavities, stronger concavities covering genito-aggenital and ano-adanal plates. Prodorsum without median crista and lateral carinae. Posterior furrows distinct and lateral rim distinct. Sigillar fields very narrow and long, median longer than laterals. Sensilli rather short with narrow pedicel and rounded head. Other setae spiniform (except vestigial exobothridial setae), short, rough. Sensilli, interlamellar and rostral setae similar in length, lamellar setae considerably shorter.

Notogaster with 15 pairs of rigid, thick, fairly short (c1

It is distinguishable from all of them by the presence of the unique tight location of genital setae g4-

6 and bifurcated setae d of femora I. It also differs from A. brasiliensis by setae h of the mentum being longer than the distance between them (versus equal in length in A. brasiliensis) and location of genital setae g6 anterior of g5 (versus posterior in A. brasiliensis); from A. iubatus by the shorter sensilli, stronger notogastral setae, and considerably shorter adanal setae ad2; from A. latebrosus by the spiniform interlamellar setae, which are different in shape than notogastral setae (versus strong setae, obtuse distally, similar to notogastral setae in A. latebrosus) and considerably shorter adanal setae ad2; from A. pervalidus by not so deeply foveolate sculpture of notogaster, shorter sensilli, stronger notogastral setae, presence of four pairs of lyrifissures (versus two pairs in A. pervalidus).

Descriptions of tritonymphs

Tritonymph 1 (Fig. 4A–D)

Identification. Most likely Indotritia (Indotritia) krakatauensis (Sellnick, 1923) General traits. Measurements. Prodorsum: length 293, width 227, height 151; notogaster: length 757, width 535, height 494. Prodorsum with setae fine and long, only exobothridial setae shorter. Notogaster with 14 pairs of setae. Ventral region. Setae h of mentum longer than distance between them. Epimeral formula: 3-1-1-1. Seven pairs of genital, two pairs of aggenital, two pairs of anal, two pairs of adanal setae. Legs. Chaetome of legs (without tarsi): I: 1-4-4(2)-5(1); II: 1-3- 3(1)-3(1); III: 2-2-2-3(1); IV: 2-2-2-1(1).

248 SYSTEMATIC & APPLIED ACAROLOGY VOL. 22 FIGURE 4. Tritonymph 1. Most likely Indotritia krakatauensis (Sellnick, 1923), tritonymph (A–C) and adult (D): A—lateral view of body; 1; B—mentum of subcapitulum; C—ventral view of opisthosoma; D—left side of ano-adanal plate of adult. Scale bars 100 μm (A, C, D), 50 μm (B).

Comparison. This tritonymph is identical to the tritonymph described by Schubart (1967). Material examined. One tritonymph: Trinidad, Chacachacare Island, Bande du Sud, 10°40'30.31"N, 61°45'26.24"W, litter and soil, 1.I.1955 (P.W. Hummelinck). One tritonymph: Trinidad, N-range, Cerro del Aripo, 10°44’N, 61°15W, litter and soil, 30.I.1955 (P.W. Hummelinck).

2017 NIEDBAŁA& ERMILOV: NEW SPECIES OF PTYCTIMOUS MITES FROM THE CARIBBEAN 249 Tritonymph 2 (Fig. 5A–D)

Identification. This tritonymph does not differ from Tritonymph 1 fundamentally. It is most likely also Indotritia (Indotritia) krakatauensis (Sellnick, 1923) or maybe Indotritia (Indotritia) bellingeri Niedbała & Schatz, 1996. General traits. Measurements. Prodorsum: length 278, width 177, height 151; notogaster: length 606, width 434, height 424. Material examined. One tritonymph: Trinidad, Gaspar Grande Island, chimney of cave, 10°40'N, 61°38'W, litter and soil, 11.I.1955 (P.W. Hummelinck).

FIGURE 5. Tritonymph 2. Most likely Indotritia (Indotritia) krakatauensis (Sellnick, 1923) or Indotritia (Indotritia) bellingeri Niedbała & Schatz, 1996; A—lateral view of prodorsum; B—lateral view of opisthosoma; C—mentum of subcapitulum; D—ventral side of opisthosoma. Scale bars 100 μm (A, B, D), 50 μm (C).

250 SYSTEMATIC & APPLIED ACAROLOGY VOL. 22 Tritonymph 3 (Fig. 6A–D)

Identification. It is most likely Arphthicarus alienus Niedbała sp. nov. General traits. Measurements. Prodorsum: length 263, width 151, height 146; notogaster: length 687, width 454, height 364. Prodorsum. Setae long and fine, interlamellar longer than lamellar and rostral setae. Notogaster with 14 pairs of setae. Ventral region. Setae h of mentum vestigial. Epimeral formula: 1-0-1-1. Seven pairs of genital, one pair of aggenital, two pairs of anal, three pairs of adanal setae. Comparison. This tritonymph is almost identical to the tritonymph of Steganacarus (Steganacarus) applicatus (Sellnick, 1920) (Niedbała 1992). Material examined. One tritonymph: Trinidad, Tetron Bay, NW Trinidad, litter and soil, 7.V.1936 (P.W. Hummelinck).

FIGURE 6. Tritonymph 3. Most likely Arphthicarus alienus Niedbała n. sp.: A—prodorsum with anterior part of notogaster, lateral view; B—lateral view of opisthosoma; C—mentum of subcapitulum; D—ventral side of opisthosoma. Scale bars 100 μm (A, B, D), 25 μm (C).

2017 NIEDBAŁA& ERMILOV: NEW SPECIES OF PTYCTIMOUS MITES FROM THE CARIBBEAN 251 Acknowledgements

We cordially thank Dr. Elizabeth A. Hugo-Coetzee (National Museum, Bloemfontein, South Africa) and two anonymous reviewers for the valuable comments, and Dr. Harry Smit (Netherlands Centre for Biodiversity Naturalis, Leiden, The Netherlands) for sending us the oribatid mite material from the Lesser Antilles and Bahamas and helping in finding the data on the localities. The present study was supported by the Russian Science Foundation (project 14-14-01134).

References

Jacot, A.P. (1924) Oribatoidea Sinensis III. Journal of the North China Branch of the Royal Asiatic Society, 55, 78–83. Mahunka, S. (1985) Neue und interessante Milben aus dem Genfer Museum LVII. Oribatida Americana 9: Antilles I (Acari). Revue Suisse de Zoologie, 92(1), 119–144. https://doi.org/10.5962/bhl.part.81605 Mahunka, S. (1998) New data on oribatids (Acari: Oribatida) from St. Lucia (Antilles). Acarologica Geneven- sia LXXXIX. Revue suisse de Zoologie, 105(4), 839–877. https://doi.org/10.5962/bhl.part.80061 Niedbała, W. (1982) Phthiracaridae (Acari, Oribatida) nouveaux d'Amerique Centrale. Redia, 65, 97–117. Niedbała, W. (1988) Phthiracaroidea (Acari, Oribatida) nouveaux du Bresil. Bulletin de la Société des Amis des Sciences et des Lettres de Poznán, Série D, sciences biologiques, 27, 111–119. Niedbała, W. (1992) Phthiracaroidea (Acari, Oribatida). Systematic Studies. Elsevier. Amsterdam-Oxford- New York-Tokyo, Pa stwowe Wydawnictwo Naukowe, 612 pp. Niedbała, W. (1994) Supplement to the classification of Phthiracaroidea, with redescriptions and descriptions of some species (Acari, Oribatida, Euptyctima). Genus, 5(1–2), 1–152. Niedbała, W. (2003) Ptyctimous mites (Acari: Oribatida) of Costa Rica. Annales Zoologici, 53(2), 259–334. Niedbała, W. (2004) Ptyctimous mites (Acari, Oribatida) of the Neotropical region. Annales Zoologici, 54(1), 1–288. Niedbała, W. (2009) New species of ptyctimous mites (Acari, Oribatida) from Cuba. Journal of Natural His- tory, 43(7–8), 423–433. https://doi.org/10.1080/00222930802610410 Niedbała, W. & Ermilov, S.G. (2015) New species and records of ptyctimous mites (Acari, Oribatida) from Cuba. Zootaxa, 4052(1), 135–142. https://doi.org/10.11646/zootaxa.4052.1.8 Niedbała, W. & Schatz, H. (1996) Euptyctimous mites (Acari, Oribatida) from the Galapagos Islands, Cocos Island, and Central America. Genus, 7(2), 239–317. Norton, R.A. & Ermilov, S.G. (2014) Catalogue and historical overview of juvenile instars of oribatid mites (Acari: Oribatida). Zootaxa, 3833, 1–132. https://doi.org/10.11646/zootaxa.3833.1.1 Schubart, H.O.R. (1967) Observations préliminaires sur la biologie d“Indotritia acanthophora” Märkel, 1964. Reviseta Brasileira de Biologia, 27(2), 165–176. Sellnick, M. (1920) Neue und seltene Oribatiden aus Deutschland. Schriften der Physikalisch-ökonomischen Gesellschaft zu Königsberg, 61/62, 35–42. Sellnick, M. (1923) Die mir bekannten Arten der Gattung Tritia Berl. Acari. Blatter für Milbenkunde, 3, 7–22.

Submitted: 2 Dec. 2016; accepted by Lizel Hugo-Coetzee: 11 Jan. 2017; published: 24 Jan. 2017

252 SYSTEMATIC & APPLIED ACAROLOGY VOL. 22